Does lemming winter grazing impact vegetation in the Canadian Arctic?

In low-productivity environments such as the tundra, it has been proposed that regular, multi-annual population cycles of lemmings could be driven by winter food depletion in years of peak abundance. If lemming population dynamics is controlled by food resources, we predict that (1) winter grazing should negatively impact the abundance of food plants, (2) this impact should be proportional to lemming density and (3) high lemming winter grazing pressure should result in reduced plant growth during the following summer. We tested these predictions on Bylot Island, Nunavut, Canada, where two species of lemmings are present: the brown (Lemmus trimucronatus) and collared lemming (Dicrostonyx groenlandicus). We installed 16 exclosures in their preferred wintering habitat (snowbeds) and annually sampled plant biomass inside and outside exclosures at snow melt and at peak growth during the summers of 2009–2012, covering a full population cycle. Winter grazing had no impact on total vascular plant or moss biomass at snow melt in all years. Among plant families, only Caryophyllaceae, which was uncommon, showed a decline. In moss taxa, a negative effect was found on Polytrichum in only 1 year out of three. Overall, plant regrowth during the subsequent summer showed annual variation and tended to be reduced in the 2 years of high lemming abundance. However, this could be a consequence of summer grazing. Overall, the impact of lemming winter grazing on plants was weak and short-lived, even in years of high lemming abundance. Therefore, our results are not consistent with the hypothesis that food depletion during winter was the cause of the lemming decline following peak abundance at our study site. Other factors may limit lemming populations and prevent them from reaching densities high enough to exhaust their food resources.     

Landscapes of fear or competition? Predation did not alter habitat choice by Arctic rodents

In systems where predation plays a key role in the dynamics of prey populations, such as in Arctic rodents, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, habitat selection by collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings depends on intra- and inter-specific densities, and there has been little agreement on the respective influences of food abundance, predators, and competition for habitat on lemming dynamics. Thus, we investigated whether predation affected selection of sedge-meadow versus upland tundra by collared lemmings in the central Canadian Arctic. We first controlled for the effects of competition on lemming habitat selection. We then searched for an additional signal of predation by comparing habitat selection patterns between 12 control plots and one large grid where lemmings were protected from predators by fencing in 1996 and 1997, but not during 5 subsequent years when we monitored habitat use in the grid as well as in the control plots. Dicrostonyx used upland preferentially over meadows and was more numerous in 1996 and 2011 than in other sample years. Lemmus was also more abundant in 1996 than in subsequent years, but its abundance was too low in the exclosure to assess whether exclusion of predators influenced its habitat selection. Contrary to the effects of competition, predation had a negligible impact on the spatial dynamics of Dicrostonyx, at least during summer. These results suggest that any differences in predation risk between the two habitats have little direct influence on the temporal dynamics of Dicrostonyx even if induced through predator–prey cycles.     

Population cycles of lemmings near Barrow,Alaska: a historical review

Current hypotheses regarding the causes of population cycling of brown lemmingsLemmus trimucronatus (Richardson, 1828), developed during long-term studies from 1950–1974. We maintain that three factors largely determine the timing and amplitude of population cycles in brown lemmings. First, a basic interaction between lemmings and vegetation sets the stage because dense populations of lemmings severely damage the vegetation, at which point lemming populations decline and remain low until the vegetation recovers. Second, opportunistic predators, mainly jaegers and owls, assemble as the snow melts during peak years and drive already declining populations to extremely low densities. Weasels are effective predators under the snow, but they appear irregularly. If weasel populations increase early in the cycle, lemming populations that normally increase dramatically during the winter in a peak year can be decimated before the snow melts. Finally, both wet summers that result in extensive flooding of the preferred habits of lemmings and freezing rains or winter thaws that cause ice formation reduce food availability, disrupt the cycle and extend the period between peak densities. Numerous observations, experimental results and simulation models support these views. Similar results regarding the importance of the available food supply, particularly in winter, and of the predation regime also have been reported for arvicoline populations at lower latitudes.     

Demography of two lemming species on Bylot Island, Nunavut, Canada

Lemmings play a key role in the tundra food web and their widely reported cyclic oscillations in abundance may have a strong effect on other components of the ecosystem. We documented seasonal and annual variations in population density, reproductive activity, survival, and body mass of two sympatric species, the brown (Lemmus trimucronatus) and collared lemmings (Dicrostonyx groenlandicus), over a 2-year period on Bylot Island, Nunavut, Canada. We live trapped and marked lemmings on two grids throughout the summer and we estimated demographic parameters using three different capture–recapture methods. All three methods are based on robust estimators and they yielded similar population density estimates. The density of brown lemmings declined markedly between the 2 years whereas that of collared lemmings was relatively constant. For brown lemmings, 2004 was a peak year in their cycle and 2005 a decline phase. Density of brown lemmings also decreased during the summer, but not that of collared lemmings. The recruitment of juvenile brown lemmings in the population increased during the summer and was higher in the peak year than in the year after, but no change was detected in collared lemmings. Survival rates of both species tended to be lower during the peak year than in the following year and body mass of brown lemmings was higher in the peak year than in the following year. We conclude that both changes in adult survival and juvenile recruitment occur during the population decline of brown lemmings.     

Habitat selection,reproduction and predation of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.     

Habitat selection and the scale of ghostly coexistence among Arctic rodents

Competition between coexisting species existing near their stable equilibrium can be obscured if they occupy separate habitats. Theories of habitat selection promise an ability to reveal the underlying ghost of competition by using isodars to infer the behavioural map of habitat selection. We tested the theory with two years of data on abundance and habitat preference by three Arctic rodent species living at low density along a gradient of wet to dry tundra on Herschel Island in Canada's western Arctic. Generalist brown lemmings exhibited a constant partial preference toward wet tundra whereas specialist collared lemmings and voles occupied the driest and wettest zones respectively. Although both lemming species compete for habitats elsewhere in the Canadian Arctic, isodar analyses suggest that the three species occupy wet and dry habitats independently of one another on Herschel Island. Competition at this large scale may be hidden at low densities, however, if the wet‐dry dichotomy is too coarse. Analyses at a finer subdivision of habitat revealed that these species coexist by using different microhabitats. Collared lemmings shifted their niche towards even drier habitat as the abundance of brown lemmings increased. We were thus able to reveal the ghost of competition lurking at large scales through a more refined analysis at smaller scales of density‐dependent habitat use.     

Lemming winter habitat choice: a snow-fencing experiment

The insulative value of early and deep winter snow is thought to enhance winter reproduction and survival by arctic lemmings (Lemmus and Dicrostonyx spp). This leads to the general hypothesis that landscapes with persistently low lemming population densities, or low amplitude population fluctuations, have a low proportion of the land base with deep snow. We experimentally tested a component of this hypothesis, that snow depth influences habitat choice, at three Canadian Arctic sites: Bylot Island, Nunavut; Herschel Island, Yukon; Komakuk Beach, Yukon. We used snow fencing to enhance snow depth on 9-ha tundra habitats, and measured the intensity of winter use of these and control areas by counting rodent winter nests in spring. At all three sites, the density of winter nests increased in treated areas compared to control areas after the treatment, and remained higher on treated areas during the treatment. The treatment was relaxed at one site, and winter nest density returned to pre-treatment levels. The rodents’ proportional use of treated areas compared to adjacent control areas increased and remained higher during the treatment. At two of three sites, lemmings and voles showed significant attraction to the areas of deepest snow accumulation closest to the fences. The strength of the treatment effect appeared to depend on how quickly the ground level temperature regime became stable in autumn, coincident with snow depths near the hiemal threshold. Our results provide strong support for the hypothesis that snow depth is a primary determinant of winter habitat choice by tundra lemmings and voles.     

Demographic response of tundra small mammals to a snow fencing experiment

Snow cover is a key environmental component for tundra wildlife that will be affected by climate change. Change to the snow cover may affect the population dynamics of high‐latitude small mammals, which are active throughout the winter and reproduce under the snow. We experimentally tested the hypotheses that a deeper snow cover would enhance the densities and winter reproductive rates of small mammals, but that predation by mustelids could be higher in areas of increased small mammal density. We enhanced snow cover by setting out snow fences at three sites in the Canadian Arctic (Bylot Island, Nunavut, and Herschel Island and Komakuk Beach, Yukon) over periods ranging from one to four years. Densities of winter nests were higher where snow depth was increased but spring lemming densities did not increase on the experimental areas. Lemmings probably moved from areas of deep snow, their preferred winter habitat, to summer habitat during snow melt once the advantages associated with deep snow were gone. Our treatment had no effect on signs of reproduction in winter nests, proportion of lactating females in spring, or the proportion of juveniles caught in spring, which suggests that deep snow did not enhance reproduction. Results on predation were inconsistent across sites as predation by weasels was higher on the experimental area at one site but lower at two others and was not higher in areas of winter nest aggregations. Although this experiment provided us with several new insights about the impact of snow cover on the population dynamics of tundra small mammals, it also illustrates the challenges and difficulties associated with large‐scale experiments aimed at manipulating a critical climatic factor.     

Comparing the genetic population structure of two species of arctic lemmings: more local differentiation in Lemmus trimucronatus than in Dicrostonyx groenlandicus

Collared and brown lemmings ( Dicrostonyx groenlandicus and Lemmus trimucronatus ) are two largely sympatric and ecologically comparable species of arctic microtine rodents, differing however in some respects which allow us to hypothesise differences in the genetic structure of their populations. Collared lemmings are particularly well adapted to life at high latitude, they occasionally emerge to the surface of the snow and may disperse over larger distances than brown lemmings – possibly even over snow and ice. This should result in more local differentiation among populations of brown lemmings than among populations of collared lemmings. We compared the genetic population structure between the two lemming species in a fragmented landscape with small islands in the central Canadian Arctic using four microsatellite loci and partial mitochondrial control region sequences. Both types of genetic markers showed higher differentiation ( F _ST values) among local populations for brown lemmings than for collared lemmings. We discuss to what extent the observed genetic differences may be explained by differences in dispersal rates in addition to differences in average effective population size.     

Positive feedback between ecological and reproductive character displacement in a young avian hybrid zone

Character displacement can reduce costly interspecific interactions between young species. We investigated the mechanisms behind divergence in three key traits-breeding habitat choice, timing of breeding, and plumage coloration-in Ficedula flycatchers. We found that male pied flycatchers became expelled from the preferred deciduous habitat into mixed forest as the superior competitor, collared flycatchers, increased in numbers. The peak in food abundance differs between habitats, and the spatial segregation was paralleled by an increased divergence in timing of breeding between the two species. Male pied flycatchers vary from brown to black with brown coloration being more frequent in sympatry with collared flycatchers, a pattern often proposed to result from selection against hybridization, that is, reinforcement. In contrast to this view, we show that brown male pied flycatchers more often hybridize than black males. Male pied flycatcher plumage coloration influenced the territory obtained in areas of co-occurrence with collared flycatchers, and brown male pied flycatchers experienced higher relative fitness than black males when faced with heterospecific competition. We suggest that allopatric divergence in resource defense ability causes a feedback loop at secondary contact where male pied flycatchers with the most divergent strategy compared to collared flycatchers are favored by selection.     

Wintering space use and site fidelity in a nomadic species,the snowy owl

Migratory species can exploit many habitats over vast geographic areas and adopt various patterns of space and habitat use throughout their annual cycle. In nomadic species, determinants of habitat use during the non‐breeding season are poorly known due to the unpredictability of their movement patterns. Here, we analysed variability in wintering space and habitat use by a highly nomadic species, the snowy owl, in eastern North America. Using 21 females tracked by satellite telemetry between 2007 and 2016, we 1) assessed how space use patterns in winter varied according to the type of environment (marine vs terrestrial), latitudinal zone (Arctic vs temperate), local snow conditions and lemming densities and 2) investigated winter habitat and site fidelity. Our results confirmed a high inter‐individual variation in patterns of habitat use by wintering snowy owls. Highly‐used areas were concentrated in the Arctic and in the marine and coastal environments. Owls wintering in the marine environment travelled over longer distances during the winter, had larger home ranges and these were divided in more smaller zones than individuals in terrestrial environments. Wintering home range sizes decreased with high winter lemming densities, use of the marine environment increased following high summer lemming densities, and a thick snow cover in autumn led to later settlement on the wintering ground. Contrary to expectations, snowy owls tended to make greater use of the marine environment when snow cover was thin. Snowy owls were highly consistent in their use of a given wintering environment and a specific latitudinal zone between years, but demonstrated flexibility in their space use and a modest site fidelity. The snowy owls’ consistency in wintering habitat use may provide them with advantages in terms of experience but their mobility and flexibility may help them to cope with changing environmental conditions at fine spatial scale.     

Spatial and temporal scaling in habitat utilization by klipspringers (Oreotragus oreotragus) determined using giving‐up densities

An animal's pattern of habitat use can reveal how different parts of its environment vary in quality based on the costs (such as predation risk) and benefits (such as food intake) of using each habitat. We studied klipspringer habitat use in Augrabies Falls National Park, South Africa using giving‐up densities (GUDs; the amount of food remaining in a resource patch following exploitation) in experimental food patches. We tested hypotheses related to how salient habitat variables might influence klipspringers' perceptions of foraging costs. At small spatial scales (3–4 m), klipspringer GUDs did not vary with cover and open microhabitats, or with the four cardinal aspects (shading) around shrubs. Adding water adjacent to food patches did not influence GUDs, showing that water is not a limiting complementary resource to food. Generally, klipspringers do not appear to be physiologically constrained. There was no difference in GUDs between four daily time periods, or between summer and winter; however, a significant interaction effect of time‐of‐day with season resulted from GUDs during the midday time period in winter being significantly higher (perceived value lower) than during the same time period in summer. At moderate spatial scales (10–60 m), klipspringer GUDs increased with distance from rocks because of increased predation risk. Based on GUDs collected at the largest scale (two 4.41‐ha grids), klipspringers preferred foraging at greater distances from drainage lines and on pebble and cobble substrates. Overall, this study has shown the efficacy of measuring GUDs to determine klipspringers' habitat utilization while foraging.     

Photoperiod differentially affects immune function and reproduction in collared lemmings (Dicrostonyx groenlandicus)

Many nontropical rodent species experience predictable annual variation in resource availability and environmental conditions. Individuals of many animal species engage in energetically expensive processes such as breeding during the spring and summer but bias investment toward processes that promote survival such as immune function during the winter. Generally, the suite of responses associated with the changing seasons can be induced by manipulating day length (photoperiod). Collared lemmings (Dicrostonyx groenlandicus) are arvicoline rodents that inhabit parts of northern Canada and Greenland. Despite the extreme conditions of winter in their native habitat, these lemmings routinely breed during the winter. In the laboratory, collared lemmings have divergent responses to photoperiod relative to other seasonally breeding rodents; short day lengths can stimulate, rather than inhibit, the reproductive system. Male and female collared lemmings were maintained for 11 weeks in 1 of 3 photoperiods (LD 22:2, LD 16:8, or LD 8:16) that induce markedly different phenotypes. Following photoperiod treatment, cell-mediated immune function as assessed by delayed-type hypersensitivity reactions was elevated in lemmings housed in LD 16:8 and LD 8:16 relative to LD 22:2. However, antibody production to a novel antigen was unaffected by photoperiod. Exposure to LD 8:16 induced weight gain, molt to a winter pelage, and in contrast to previous studies, regression of the male, but not the female, reproductive tract. In conclusion, these data indicate that components of immune function among collared lemmings are responsive to changes in day length.     

Highly Overlapping Winter Diet in Two Sympatric Lemming Species Revealed by DNA Metabarcoding

Sympatric species are expected to minimize competition by partitioning resources, especially when these are limited. Herbivores inhabiting the High Arctic in winter are a prime example of a situation where food availability is anticipated to be low, and thus reduced diet overlap is expected. We present here the first assessment of diet overlap of high arctic lemmings during winter based on DNA metabarcoding of feces. In contrast to previous analyses based on microhistology, we found that the diets of both collared (Dicrostonyx groenlandicus) and brown lemmings (Lemmus trimucronatus) on Bylot Island were dominated by Salix while mosses, which were significantly consumed only by the brown lemming, were a relatively minor food item. The most abundant plant taxon, Cassiope tetragona, which alone composes more than 50% of the available plant biomass, was not detected in feces and can thus be considered to be non-food. Most plant taxa that were identified as food items were consumed in proportion to their availability and none were clearly selected for. The resulting high diet overlap, together with a lack of habitat segregation, indicates a high potential for resource competition between the two lemming species. However, Salix is abundant in the winter habitats of lemmings on Bylot Island and the non-Salix portion of the diets differed between the two species. Also, lemming grazing impact on vegetation during winter in the study area is negligible. Hence, it seems likely that the high potential for resource competition predicted between these two species did not translate into actual competition. This illustrates that even in environments with low primary productivity food resources do not necessarily generate strong competition among herbivores.     

Population dynamics of the collared lemming and the tundra vole at Pearce Point,Northwest Territories,Canada

From 1987 to 1989 we monitored population changes during summer of the collared lemming (Dicrostonyx groenlandicus) and the tundra vole (Microtus oeconomus) at Pearce Point, Northwest Territories, Canada (69° 48 N, 122° 40 W). Populations on four study areas did not cycle but remained at low density (<3/ha) each year and continued at low numbers for the following 3 years (Reid et al. 1995). Lemming numbers often declined throghout the summer in spite of continous reproduction, and population recovery occurred overwinter. Heavy predation losses of radio-collared lemmings occurred each summer and this lemming population may be trapped in a predator-pit. Collared lemmings breed in winter and only because of winter population growth do these populations persist. Tundra vole numbers increased rapidly in most summers but usually declined overwinter. Tundra voles do not seem able to sustain winter reproduction in this extreme environment and this prevents them from reaching high density because of the short summer. Population growth in both these rodents could be prevented by poor food or by predation losses, and landscape patchiness may also help to prevent population growth. For lemmings we do not think that a shortage of shelter or intrinsic limitations could be restricting population increase at Pearce Point. This is the first detailed study of a non-cyclic collared lemming population.     

Photoperiodic effects on post-weaning growth and food consumption in the collared lemming Dicrostonyx groenlandicus

The effects of long 'summer' (22L:2D) and short 'winter' (2L:22D) photoperiod on post-weaning growth and food consumption of the collared lemming were examined. Growth was described by Gompertz equations. After 10 weeks, lemmings maintained under short photoperiods developed white winter pelage and heavy bifid claws; body mass and estimated asymptotic mass were significantly greater than for lemmings reared under long photoperiod. Sexual dimorphism in body size was observed within treatments, males growing larger than females. There were no significant differences in the overall growth rate constant k as a result of either sex or photoperiod treatment. However, instantaneous growth rates (dW/dt) were significantly higher in 'winter' lemmings. Mass at weaning was a significant determinant of adult mass at 90 days.
Cumulative food consumption at 90 days was not significantly different between photoperiod groups, even though 'winter' lemmings gained more than double the mass of their 'summer' counterparts over the post-weaning period. Increased gut length observed for winter morphs may act as the physiological mechanism promoting greater gross energy efficiency.
The present study suggests that the overwintering strategy of the collared lemming with respect to body mass changes and energy requirements differs considerably from that of other microtines.     

Summer grazing by voles and lemmings upon subarctic snow-bed and tall herb meadow vegetation – an enclosure experiment

Ther hypothesis of the exploitation of ecosystems predicts that plant communities in productive habitats should experience a low grazing pressure, while communities in less productive habitats should be structured by intense grazing. Thus, if a high population density of rodents was to be established in these communities and they could graze without predation, the plant community in the productive habitat should be more affected than the community in the less productive habitat. This was tested by building enclosures for grey-sided voles Clethrionomys rufocanus and lemmings Lemmus lemmus in a productive tall herb meadow and a less productive snow-bed on Finnmarksvidda, northern Norway. The simulated rodent population densities varied from 150 to 750 ind. ha^-1, and the rodents were allowed to graze for 18 to 55 d. The shoot mortality of relatively abundant plant species were monitored together with plant community structure, and the above-ground biomass were harvested at the end of the experiment. The shoot mortality were generally low, the significantly grazed species were either known preferred species or species with a grazing-sensitive morphology. The community structure did not show any major changes. The aboveground biomass of woody plants was significantly lower in the enclosures on the snow-bed, but not in the tall herb meadow. The total biomass did not differ significantly. These results are somewhat ambiguous with reference to the predictions from the food chain hypothesis, and the conclusion were drawn that intense grazing by voles and lemmings during 18 to 55 d during the growing season does not seem to be sufficient to greatly influence shoot mortality and structure of the studied plant communities.     

Resource partitioning by mammalian herbivores in the high Arctic

Willow (Salix arctica) and sedges (Carex stans and Eriophorum triste) were the dominant plants available as forage for herbivores in the high Arctic of Greenland. Willow leaves were of high quality as forage in early stages, of phenology, but crude protein and digestibility declined markedly by late stages whereas sedges, remained high in forage quality throughout the growing season. Densities of fecal pellets indicated that muskoxen (Ovibos moschatus) made heaviest use of sedge-dominated vegetation types in both winter and summer, although increased use of willow communities was observed in early summer. Hares (Lepus arcticus) favored willow-dominated communities in both winter and summer. Evidence of collared lemming (Dicrostonyx groenlandicus) winter use was mainly in willow-dominated communities where snow had accumulated, whereas in summer they were present in drier habitas dominated by willows, but with greater plant diversity. Analyses of plant tissues in feces indicated that graminoids composed over 60% of the diet of muskoxen in winter and over 40% in summer. Willows were of nearly equal importance in the muskox diet in summer, and forbs, Dryas integrifolia, and moss collectively composed over 20% of the diet in both summer and winter. Grass accounted for nearly 50% of the diet of hares in both summer and winter, with willows, forbs, and moss accounting for most of the remainder. Willows and graminoids dominated the diet of lemmings, with willows being somewhat more important in summer and graminoids in winter. Moss was a noteworthy dietary component of lemmings. Differences in body and digestive-tract morphology among the three mammalian herbivores account for differences in locomotive efficiency, predator avoidance, and foraging efficiency which interact with vegetation quality, density, and patchiness. The resulting patterns of use of the landscape result in minimal overlap in use of forage resources and help to explain the distribution and co-existence of high Arctic herbivores.     

Density‐dependent habitat selection predicts fitness and abundance in a small lizard

Density‐dependent habitat selection has been used to predict and explain patterns of abundance of species between habitats. Thermal quality, a density‐independent component of habitat suitability, is often the most important factor for habitat selection in ectotherms which comprise the vast majority of animal species. Ectotherms may reach high densities such that individual fitness is reduced in a habitat due to increased competition for finite resources. Therefore, density and thermal quality may present conflicting information about which habitat will provide the highest fitness reward and ectotherm habitat selection may be density‐independent. Using ornate tree lizards Urosaurus ornatus at 10 sites each straddling two adjacent habitats (wash and upland), we tested the hypothesis that habitat selection is density‐dependent even when thermal quality differs between habitats. We first tested that fitness proxies decline with density in each habitat, indicating density‐dependent effects on habitat suitability. We also confirmed that the two habitats vary in suitability (quantified by food abundance and thermal quality). Next, we tested the predictions that habitat selection depends on density with isodar analyses and that fitness proxies are equal in the two habitats within a site. We found that monthly survival rates decreased with density, and that the wash habitat had more prey and higher thermal quality than the upland habitat. Lizards preferred the habitat with more food and higher thermal quality, lizard densities in the two habitats were positively correlated, and fitness proxies of lizards did not differ between habitats. These patterns are consistent with density‐dependent habitat selection, despite differences in thermal quality between habitats. We expect that density‐dependent habitat selection is widespread in terrestrial ectotherms when densities are high and temperatures are close to their optimal performance range. In areas where thermal quality is low, however, we expect that depletable resources, such as food, become less limiting because assimilating resources is more difficult.     

Migratory strategies, fawn recruitment, and winter habitat use by urban and rural mule deer (Odocoileus hemionus)

We examined the migratory strategies of mule deer (Odocoileus hemionus) using adjacent urban and rural winter ranges in northern Utah in relation to deer demography and patterns of habitat use. Urban deer were more likely to be migratory than rural deer, even though migratory animals from the two herds intermixed on a common, high-elevation summer range. Urban deer exhibited lower fawn recruitment than rural deer; but within each herd, demographic characteristics of migratory and nonmigratory animals suggested that game theory explained the ratios of deer adopting each behavior. Estimates of animal numbers and available habitat did not reveal clearly whether deer densities differed on the two winter ranges. However, patterns of habitat use by urban deer were so clustered around areas of concealment vegetation that animals probably experienced higher local densities than rural animals. In addition, the clustered patterns of habitat use by urban deer resulted in incomplete use of available forage. This may have contributed to the relatively poor fawn recruitment by urban deer, a phenomenon that appeared to be perpetuated by their strong fidelity to winter range.